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Copyright: 2014 Vogel et al. This is an open-access article distributed under the terms
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On Trimeresurus sumatranus (Raffles, 1822), with the designation of a neotype and the description of a new species of
pitviper from Sumatra (Squamata: Viperidae: Crotalinae)
1
Society for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, GERMANY 2Reptiles & Amphibiens, UMR 7205 OSEB, Dpartement
Systmatique et volution, CP 30, Musum National dHistoire Naturelle, 57 rue Cuvier, 75231 Paris Cedex 05, FRANCE 3Laboratory of
Herpetology, Museum Zoologicum Bogoriense, Zoology Division, Research Center for Biology, Indonesian Institute of Sciences, Widyasatwa Loka
Jl. Raya Jakarta-Bogor Km 46, Cibinong 16911, INDONESIA
1
Introduction
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Vogel et al.
Boulengers appraisal was followed by subsequent workers, until the works of van Lidth de Jeude (1922) and
especially Brongersma (1933). This latter author was the
first to show that T. sumatranus and T. hageni were valid,
distinct species, and he provided new characters separating both species from one another.
Furthermore, it is worth noting that Schlegel (1826)
described the taxon Cophias wagleri (nec Cophias wagleri Boie, 1827) as a replacement name for Coluber
sumatranus (Raffles, 1822). We refer to Savage et al.
(2012) for a discussion on the early confusion between
these species. Nevertheless, other authors such as Schlegel (1837), Gray (1842: 48; 1849: 10), Cantor (1847:
1042, Pl. 40: Fig. 9), Gnther (1858: 266), Peters (1862:
671) and later as Ouwens (1916: Pl. 15: Fig. 22 and 22a)
also confused in part or totally Coluber sumatranus with
Cophias wagleri Boie, 1827, a totally different species
now known as Tropidolaemus wagleri.
Loveridge (1938: 45) described Trimeresurus sumatranus malcolmi (Type locality: Sungii River, near
Bunduntuan, Mount Kinabalu, British Nord Borneo,
a river in the vicinity of Bundu Tuhan, on the southern
slopes of Mt. Kinabalu, state of Sabah, Borneo, Federation of Malaysia). This subspecies was regarded as valid
by all subsequent authors for some populations of northern Borneo whereas other populations of this island were
referred to the nominative subspecies (David and Ineich
1999; McDiarmid et al. 1999; Malkmus et al. 2002).
Subsequently, Trimeresurus sumatranus malcolmi was
raised to full species status by Stuebing and Inger (1998).
As a consequence, Trimeresurus sumatranus was subsequently considered monotypic.
More recently, two revisions of the systematics of
these two species were published by Sanders et al. (2002,
2004). Results of the first publication, which was based
on scalation characters, pattern and coloration, can be
summarized as follows: (1) T. sumatranus and T. hageni
are clearly separate as shown by canonical multivariate analysis; (2) T. sumatranus inhabits South Thailand,
Borneo, and central western Sumatra, whereas T. hageni
is living in North and South Sumatra, Thailand, Malaysia, Singapore, Nias, and Siberut; (3) the authors showed
clear differences between populations of T. sumatranus
inhabiting the central part of western Sumatra and that
one living on Borneo; and (4) morphological differences,
especially in males, were pointed out between populations of the islands of Nias and Siberut on the one hand,
and all other populations on the other hand. In these islands, specimens referable to Trimeresurus hageni show
some characters of the pattern typical to T. sumatranus,
such as black dorsal crossbars and the presence of dark
edges on head scales. This partial similarity has led to
erroneous records of T. sumatranus from these islands.
In contrast, Sanders et al. (2004) included molecular analyses and considered all species of the subgenus
Parias Gray, 1849 as defined by Malhotra and Thorpe
(2004) (as a genus). For the species treated here, the
Amphib. Reptile Conserv. | amphibian-reptile-conservation.org
Table 1. List of morphological characters and variables used in this study and their abbreviations.
Number
Abbreviation
Character
SVL
Snout-vent length
TaL
Tail length
TL
Total length
TaL/TL
Morphometry
Scalation
5
6
Ven
Ventral plates
Sc
Subcaudal plates
10
Cep
11
InN
Internasal scale(s)
12
InN sep
13
14
Supralabial scales
15
16
17
18
CtotSL
19
IL
Infralabials
Pattern
20
21
22
23
24
25
26
27
28
Color of eyes
29
30
Vogel et al.
The analyses of external morphological data were
based on comparisons of statistical values (mean value
and standard deviation). A test of Mann-Whitney (U
test; see Siegel 1956) was applied as necessary. Calculations were run online on the website: http://elegans.som.
vcu.edu/~leon/stats/utest.html (last accessed on 14 July
2014). Abbreviations are: n: number of specimens; x :
mean value; s: standard deviation; P: probability of occurrence of a value as extreme as or more extreme than
the observed value; U: the statistic in the Mann-Whitney
test.
The color of the eyes is shown here to be a taxonomic
character. However, it is problematic as it cannot be observed in preserved specimens. According to our observations, the eye color in adult animals is stable for each
species and sex (Vogel et al. 2004). In the species treated
here, there was no sexual dimorphism in eye coloration.
The color of the tail is diagnostic and we recognize two
patterns: uniform reddish-brown with dark margins, or
mottled, for specimens with a mixture of brown and
green colors on the tail.
Results
In our sample of 53 specimens referred to Trimeresurus sumatranus, as currently defined, we noticed that
nine specimens from western Sumatra differed in several morphological characters from other populations.
Trimeresurus sumatranus (Raffles, 1822) was briefly described (as Coluber sumatranus) without any designation
of a name-bearing type. Therefore, we first redefine this
species and note intraspecific variation of the characters
examined. We then designate a neotype for this species
in agreement with Art. 75.3.1 to 75.3.7 of the International Code of Zoological Nomenclature (I.C.Z.N. 1999;
merely designated below as the Code).
Museum abbreviations
Indonesia
Other abbreviations
Sumatra. Bengkulu Province. MZB 1035, Gunung Gedang; MZB 2180, Muara Aman, North Bengkulu;
MZB 3718, ZMB 661778; ZMB 76340, ZMB 70490,
Bengkulu; ZMH R06936, Lebong-Tandai (301S
1015E). Jambi Province. MZB 457, Jambi. Lampung
Province. MZB 2166, Rimba; MZB 2219, Propinsi
Lampung. Sumatera Barat Province. OMNH R21356,
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Federation of Malaysia
Borneo (East Malaysia). State of Sabah. FMNH
23994952, FMNH 2399578, Tenom District; FMNH
239959, Sipitang District. State of Sarawak. BMNH
91.8.29.33, Mt. Dulit, Miri District, Miri Division;
BMNH 1978.1879, Gunung Mulu National Park, Miri
District, Miri Division; FMNH 1386878, FMNH
138690, FMNH 148829, Kapit District, Kapit Division;
FMNH 158671, Bintulu, Bintulu Division.
West Malaysia. State of Johore. BMNH 1971.1532, Panti Forest Reserve, South Johore. State of Pahang. ZRC
2.2929, Kuala Tahan; ZMB 69982, Pahang. State of
Trengganu. BMNH 1974.50013, Gunong Lawit.
Thailand
Yala Province. BMNH 1936.9.12.3, Betong, Pattani.
Vogel et al.
Fig. 1 A-C. Trimeresurus sumatranus, ZFMK 76340, neotype of Coluber sumatranus Raffles, 1822, southwest Sumatra, Bengkulu
Province, Sumatra. A. dorsal view of the body, B. ventral view of the body, C. lateral view oft the head. Photo: G. Vogel.
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C
Fig. 2 A-C. Trimeresurus sumatranus, RMNH 1583, Holotype of Trigonocephalus
formosus Mller and Schlegel, 1842, from Padang, Sumatera Barat Province, Sumatra A. dorsal view of the
body, B. ventral view of the
body, C. lateral view of the
head. Photo: G. Vogel.
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Vogel et al.
Fig. 3A. MZB.Ophi.5452 live holotype of Trimeresurus gunaleni spec. nov. from Mt. Sibayak, ca. 1,800 m a.s.l., west of Brastagi,
Sumatera Utara Province, Sumatra, adult female. Photo: G. Vogel.
Fig. 3B. MZB.Ophi.5452 live holotype of Trimeresurus gunaleni spec. nov. from Mt. Sibayak, ca. 1,800 m a.s.l., west of Brastagi,
Sumatera Utara Province, Sumatra, adult female. Photo: G. Vogel.
Amphib. Reptile Conserv. | amphibian-reptile-conservation.org
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Fig. 4A. Live male of Trimeresurus gunaleni spec. nov. from Mt. Singkut, 1,600 m a.s.l., Sumatra Utara Province, Sumatra. Photo:
G. Vogel.
Fig. 4B. Live male of Trimeresurus gunaleni spec. nov. from Mt. Singkut, 1,600 m a.s.l., Sumatra Utara Province, Sumatra. Photo:
G. Vogel.
Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (10)
Vogel et al.
Fig. 5. Comparative dorsal view of the head of Trimeresurus gunaleni spec. nov. (left) and T. sumatranus (right). Left from above:
Male, female (holotype), male, all from Sumatra Utara Province, Sumatra alive, right adult female alive from Bengkulu Province,
Sumatra, adult male alive from Bengkulu Province, Sumatra, preserved female from Borneo. Photos: N. Maury.
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Fig. 6. Comparative lateral view of the head of Trimeresurus gunaleni spec. nov. (left) and T. sumatranus (right). Left from above:
Male, female (holotype), male, all from Sumatra Utara Province, Sumatra alive, right adult female alive from Bengkulu Province,
Sumatra, adult male alive from Bengkulu Province, Sumatra, preserved female from Borneo. Photos: N. Maury.
Vogel et al.
Fig. 7. Comparative ventral view of the head of Trimeresurus gunaleni spec. nov. (left) and T. sumatranus (right). Left from above:
Male, female (holotype), male, all from Sumatra Utara Province, Sumatra alive, right adult female alive from Bengkulu Province,
Sumatra, adult male alive from Bengkulu Province, Sumatra, preserved female from Borneo. Photos: N. Maury.
Body scalation
Head scalation
Vogel et al.
Diagnosis
Hemipenis
Body scalation
Table 2. Main characters to distinguish between the species of the Trimeresurus sumatranus complex, source specimens from Appendix I, if not noted different.
Characters
N males/females
Middorsal scale rows
Trimeresurus
gunaleni
spec. nov.
Trimeresurus
sumatranus
Sumatra
Trimeresurus
sumatranus
Borneo
Trimeresurus
sumatranus
Peninsular Malaysia
Trimeresurus
malcolmi1
5/4
2/13
3/19
2/5
3/4
21
21
21
21
19
Ventrals males
162179
179182
182185
178183
169173
Ventrals females
164174
175186
176191
180186
168174
Subcaudals males
7172
6670
6671
6970
6481
Subcaudals females
5866
5768
5464
6166
6164
Total length
1170
1152
1350
1220
1330
0.2010.210
0.1600.166
0.1540.168
0.1500.161
0.1600.1794
0.1440.180
0.1300.159
0.1280.150
0.1340.160
0.1585
Thin
Broad
Broad
Broad
Absent
No
Yes
Yes
Yes
Yes
No
Yes
Yes
Yes
Yes
Tail reddish
No
Yes
Yes
Yes
Yes
Eye in life
Green
Dark grey
Dark grey
Dark grey
Dark grey
Head scalation
Vogel et al.
Fig. 8. Live female of Trimeresurus sumatranus from vincity of Padang Panjang, Sumatera Barat Province, Sumatra. Photo: G.
Vogel.
Distribution (Fig. 8)
Indonesia
Sumatra. Known from the provinces of Sumatera Barat,
Jambi, Bengkulu, and Lampung, in Barisan Range.
Borneo (Kalimantan). Seemingly throughout the island.
Federation of Malaysia
Borneo. Known from the States of Sabah and Sarawak.
West Malaysia. Definitely recorded from the States of
Perak (Sukumaran 2002 as Tropidolaemus wagleri, pers.
comm.), Johore, Pahang, and Trengganu.
Thailand
Recorded only from Yala Province.
In contrast to Sanders et al. (2004), we confirm the occurrence of T. sumatranus in extreme southern Thailand and
West Malaysia. Examined specimens present the combination of all scalation and pattern characters, both of
the head and body, in full agreement with the definition
of this species. They all differ from Trimeresurus hageni
and we could not find any reason for not referring them
to T. sumatranus. The range of T. sumatranus in Sumatra
is wider than indicated in Sanders et al. (2004) but the
records from the Indonesian islands of Bangka, Belitung,
Nias, Simeulue, and the Mentawai Archipelago (see, for
example, Brongersma 1933; Dring et al. 1990), are now
referred to the T. hageni group.
Natural History
This beautiful species inhabits regions typically covered
with equatorial rainforests, lowland tropical wet forests,
and tropical wet submontane forests, from sea level up
to about 1,000 m. The species shows a predilection for
lowlands in Borneo but, seemingly, only for hilly areas
at elevations between 650 and about 900 m in Sumatra
(Ryabov et al. 2002; Gumprecht et al. 2003; Sanders et
al. 2004). This pitviper is found in tropical forests, along
clearings, in bamboo thickets, mangroves, swamps,
plantations, and cultivated fields such as coffee and tea
estates. However, in Sumatra, all specimens recorded
by Ryabov et al. (2002) and Gumprecht et al. (2003) in
Bengkulu Province (Sumatra) were found in forest, none
in cultivated areas or near villages.
Fig. 9. Live female of Trimeresurus sumatranus from Bengkulu Province, Sumatra. Photo: G. Vogel.
Fig. 10. Live female of Trimeresurus malcolmi from Mount Kinabalu, Sabah, Borneo. Photo: M. Dehling.
Vogel et al.
Fig. 11. Live female of Trimeresurus malcolmi from Mount Kinabalu, Sabah, Borneo. Photo: M. Dehling.
Fig. 12. Live male of Trimeresurus toba from vincity of Padang Panjang, Sumatera Barat Province, Sumatra, a species sympatric
with T. gunaleni spec. nov. Photo: G. Vogel.
Fig. 13. MZB.Ophi.5452 holotype of Trimeresurus gunaleni spec. nov., adult female. Photo: N. Maury.
Fig. 14. Live female of Trimeresurus sumatranus from Bengkulu Province, Sumatra. Photo: N. Maury.
Vogel et al.
Trimeresurus sumatranus often occurs along the
banks of rivers, ponds, and other watered areas. This diurnal and nocturnal species is chiefly arboreal but lives
in the lower vegetation such as in thick bushes, shrubs,
and the tangled lower tree foliage up to 2.5 m above the
ground, where it proves to be a skilled climber. Ryabov et
al. (2002) found specimens basking in early morning. Trimeresurus sumatranus feeds on small mammals, frogs,
lizards, and frogs. It is oviparous, but little is known on
its breeding habits. Ryabov et al. (2002) mentioned a
clutch of 17 eggs that were guarded by the female; we
refer to Ryabov et al. (2002) and Gumprecht et al. (2003)
for additional data on the biology of this species.
In our sample of specimens identified in collections
as Trimeresurus sumatranus, we identified a total of six
specimens that present noteworthy morphological differences with the species as defined above. We also noted
the same differences in three specimens that were kept
alive. As these differences with T. sumatranus are constant, we consider these specimens to be referable to a
distinct species that we here describe as:
Trimeresurus sumatranus (nec Coluber sumatranus Raffles, 1822): Sanders et al. (2002: 107, part.; 2004: 722,
part.).
Holotype
MZB.Ophi.5452, adult female, from Mt. Sibayak, ca.
1,5002,200 m a.s.l., west of Brastagi (Berastagi), Karo
Regency (Kabupaten Karo), Sumatera Utara Province,
Sumatra, Indonesia. Collected by the team of Danny Gunalen, Hidekazu Miyake, Cho Sangyeon, and Moon Suk
Cha.
Diagnosis
coloration with interstitial skin forming irregular, hollow, black dorsal crossbands, with a thin, pale ventrolateral line; (3) 21 DSR at midbody; (4) first supralabial
totally separated from nasal scale; (5) large internasals,
most usually separated by one scale, only exceptionally
in contact; (6) three supralabials, third, fourth, fifth SL in
contact with subocular; (7) supraoculars large but elongate, separated by 57 cephalic scales; (8) tail long, with
a ratio TaL/TL between 0.201 and 0.210 in males and
0.144 and 0.180 in females; (9) 162179 VEN; (10) 58
72 SC (males: 7172; females: 5866); (11) eye yellowish-green in life and preservative; (12) cephalic scales
strongly and broadly edged with black but not forming
streaks; (13) no black postocular streak; (14) venter
greenish-yellow or pale green, uniform, with posterior
margin of ventrals paler green; and (15) tail greyish-red,
rusty brown or reddish-brown, mottled with green crossbars anteriorly.
Main characters separating T. gunaleni spec. nov.
from other taxa of the complex of T. sumatranus are
summarized in Table 2. Trimeresurus gunaleni spec. nov.
mainly differs from T. sumatranus by (1) a lower number
of ventrals in males (162179, x = 168.4 vs. 178185, x
= 181.5; U = 33.5, P < 0.005) and females (164171, x
= 169.5 vs.175191, x = 183.3); (2) a higher value of the
ratio TaL/TL in males (0.2010.210, x = 0.206 vs. 0.150
0.168, x = 0.161); (3) the color of the tail with hues of
red throughout mottled with green crossbars anteriorly
vs. green as the body on its anterior half, becoming more
or less abruptly red (see above description) posteriorly,
strongly reticulate with black; (4) the color of the eyes:
green or yellowish-green in T. gunaleni spec. nov. vs.
dark brown, dark grey, or bronze in T. sumatranus; (5)
the color of the ventral scales, which are green with a
paler posterior margin in T. gunaleni spec. nov. vs. pale
green with a dark grey or black posterior margin in T.
sumatranus.
Trimeresurus gunaleni spec. nov. differs from T. malcolmi by (1) the number of dorsal scales around midbody (21 vs. 19); (2) a higher value of TaL/TL in males
(0.2010.210, x = 0.206 vs. 0.1600.179, x = 0.162, s
= 0.009); (3) the presence of a white lateral stripe in T.
gunaleni spec. nov., missing in T. malcolmi; (4) the color
of the tail: greyish-red, rusty brown or reddish-brown,
mottled with green crossbars anteriorly in T. gunaleni vs.
greenish-orange, salmon or pinkish-red, strongly reticulated with black in T. malcolmi; below, the tail is yellowish green anteriorly, turning to brown posteriorly in T.
gunaleni spec. nov. vs. green or greenish-yellow on its
anterior half, with subcaudal scales broadly edged with
black producing a conspicuous reticulation; (5) the color
of the eyes: yellowish-green in T. gunaleni spec. nov. vs.
dark grey in T. malcolmi.
Trimeresurus gunaleni spec. nov. differs from T. hageni by (1) a lower number of ventrals in males (162179,
x = 168.4 against 177189, x = 181.8; U = 139.5, P <
0.001) and females (164171, x = 169.5 vs.176196, x =
September 2014 | Volume 8 | Number 2 | e80
Etymology
Body elongate, compressed; head elongate, distinctly triangular, wide at its base, clearly distinct from the neck,
flattened anteriorly, thick posteriorly, 1.6 times as long as
wide; snout long, round when seen from above, strongly
obliquely truncated when seen from the side, with a moderate canthus rostralis, totaling 32.0 % of head length,
and 2.7 times as long as diameter of eye; a large oval nostril piercing in the middle of nasal scale; nostril-loreal pit
distance about 0.5 times the distance between the nostril
and the eye; eye average, totaling 0.65 times the distance
between the lower margin of eye and upper lip border;
tail rather long, tapering, and prehensile.
SVL 995 mm, TaL 195 mm, TL 1,170 mm; largest
head width 35.0 mm; ratio TaL / TL 0.167.
Body scalation
Head scalation
Vogel et al.
The head is deep green above (bright emerald green in
life) and on the temporal region, with scales of the snout,
preoculars, supraoculars, cephalic, occipital, and lower
temporal scales narrowly edged with black and surrounded with black interstitial skin, producing a conspicuous
pattern of a mixed black background with bright green
spots; no cephalic or occipital streaks; supraoculars narrowly edged with black; top of rostral black; anterior
supralabials greenish-yellow, distinctly paler than upper
head surface, others supralabials bright yellow in life;
firstthird SL narrowly edged with black posteriorly; no
postocular streak; upper temporals green as the upper
head surface. Chin and throat pale bluish-grey (cream in
preservative); mental and first three infralabials greenish-yellow; other infralabials more or less marbled with
greenish-yellow; posterior gular scales dotted with green.
The venter is uniformly yellowish-green, with the
posterior edge of each ventral pale bluish-grey, distinctly
paler than the background color of the venter. The tail is
greenish-yellow on the first two subcaudals then greyishred throughout as the upper surface of tail, with scales
narrowly edged with pale grey.
Hemipenis
Unknown.
Body scalation
Head scalation
As described for the holotype, with the following variation for major characters: internasals separated by one
small scale in 8/9 specimens, in contact only in specimen ZMB 29642; only two canthal scales on each side
in all specimens, not larger than adjacent snout scales or
slightly smaller, bordering the canthus rostralis between
the internasal and corresponding supraocular; two small
postoculars, in contact with first temporals or followed
by 23 small scales between postoculars and first temporals; one large, elongate, subtriangular supraocular on
each side, 1.62.1 times as long as wide, 1.01.3 times
as wide as internasal, not indented by adjacent cephalic
scales; 34 enlarged scales on upper snout surface on a
line between the scale separating the internasals and a
line connecting the anterior margins of eyes, smooth, and
juxtaposed; 57 cephalic scales (5: 1/9 specimens; 6: 5/9;
7: 3/9) on a line between supraoculars, smaller than upper snout scales, smooth, flat, and juxtaposed; occipital
scales not enlarged and smooth; temporal scales smooth,
large, subequal, in two or three rows; 810 supralabials
(8: 2/18 occurrences; 9: 12/18; 10: 4/18); third, fourth,
fifth SL in contact with subocular in all specimens; first
SL always separated from nasal; second SL tall, entirely
bordering the anterior margin of the loreal pit, always in
contact with nasal; third SL longest and highest, 1.21.5
times as long as high; fourth SL higher than long; fifth SL
tall and narrow; 1012 IL (10 or 11 in most specimens);
scales of the first pair longitudinally in contact; first three
Table 3. Morphological characters of the paratypes of Trimeresurus gunaleni spec. nov. M: male, F: female, for other abbreviations
see Table 1.
Collection number
SC
SL
SL touching
sublabial
Cep
IL
Do
162
71
9/9
11/10
21
175
inc.
9/9
11/11
21
0.205
179
inc.
9/9
11/11
21
0.144
174
58
10/10
11/11
21
182
0.158
169
58
9/10
11/11
21
88
0.180
164
66
9/9
?*/ ?*
21
Sex
SVL (mm)
TaL (mm)
TaL/ TL
VEN
NHMW 28159:1
732
195
0.210
NHMB 2599
651
inc.
inc.
ZMB 29642
638
165
NHMW 23909:4
309
52
NHMW 28159:2
972
SMF 52844
400
blue in life). The tail is greyish-red or rusty-red throughout, with scales narrowly edged with cream to pale grey,
and mottled with incomplete cream, pale grey or pale
greenish-grey rings (pale greyish-green in life), present
on the anterior half of the tail or throughout.
The head is deep green above and on the temporal
region; scales of the snout, preoculars, supraoculars,
cephalic, occipital and lower temporal scales narrowly
edged with black and entirely surrounded with interstitial
black skin, producing a conspicuous pattern made of a
black background mixed with bright green spots; no
cephalic, occipital, or postocular streaks; supraoculars
narrowly edged with black; top of rostral usually black;
anterior supralabials green or yellowish-green, distinctly
paler than upper head surface, others supralabials yellow,
greenish-yellow or green; firstthird SL usually narrowly
edged with black on their posterior edge; upper temporals green as the upper head surface. Chin and throat
cream (pale bluish-grey in life); mental and first three infralabials greenish-yellow or pale yellowish-green; other
infralabials more or less marbled with greenish-yellow;
posterior gular scales sometimes dotted with greyishgreen or green spots.
Vogel et al.
The venter is uniformly bluish-green or yellowishgreen in preservative (yellow, greenish-yellow, or green
in life), with the posterior edge of each ventral pale bluish-grey or greyish-green, distinctly paler than the background color of the venter. The tail is greenish-yellow or
green anteriorly on a distance varying from the first subcaudals to the middle of the tail then greyish-red or rustyred throughout as the upper surface of tail, with scales
narrowly edged with pale grey and with cream, pale grey
or pale greenish-grey blotches (pale greyish-green in
life) corresponding to the rings of the upper surface.
Sexual dimorphism
Distribution
Discussion
Indonesia
Sumatra. Endemic; Trimeresurus gunaleni spec. nov. is
known only from two provinces: Sumatera Barat (Solok
and Padang Mountains) and Sumatera Utara (Mt. Sibayak, Mt. Sinabung and Mt. Singkut near Berastagi).
This species can be expected in higher elevations all
over the mountainous areas of Sumatra.
Natural History
Trimeresurus gunaleni spec. nov. inhabits regions typically covered with tropical moist montane forests, from
1,500 m to as high as at least 2,000 m, perhaps as much
as 2,200 m, where it has been observed by local insect
collectors (Figs. 15 and 16). There is no record of populations lower than 1,500 m. On Mount Sibayak, Danny
Gunalen collected specimens of Trimeresurus hageni at
elevation of 500 m, and Tropidolaemus wagleri at 200 m.
Trimeresurus gunaleni is clearly isolated as a high montane dweller.
The female holotype of T. gunaleni spec. nov. was collected during the daytime in dense humid montane forest scattered with tiny springs. The snake was resting on
the ground under tree roots. In another instance, a male
was seen perched at night on a tree branch at about two
m above the ground. None of the specimens were found
Amphib. Reptile Conserv. | amphibian-reptile-conservation.org (25)
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Vogel et al.
Gernot Vogel was born in Heidelberg, Germany and received a Ph.D. in Chemistry. Dr. Vogel is now working
as a chemist at an international company, doing research and registration of plant protection products in Hirschberg, close to Heidelberg. Beginning as a reptile keeper, Dr. Vogel developed a great interest in the snake fauna
of the Oriental region. He has concentrated his work on southwestern China, Indonesia, and tropical India. Dr.
Vogel has revised large snake genera with a wide distribution, and this has been achieved through international
collaborations with, for example, the institutes of Chengdu and Kunming in China, with Lipi and KPH Salvator
in Indonesia, and with various Indian groups including ARRS and ANET. His research is based on specimen
collections all over the world and on field research in the regions cited above. Dr. Vogel has authored or coauthored the following books: The Snakes of Sumatra; The Snakes of Sulawesi; Amphibians and Reptiles of
Mount Kinabalu; and parts I to III in the Terralog Series on the venomous snakes of the world.
Patrick David (born in 1959 near Paris, France), received a Ph.D. in polymers chemistry at the University
of Paris-Orsay. Dr. David developed an early interest in herpetology as a reptile keeper and then turned to the
systematics of Asian reptiles. He is, or has been a member of several international herpetological societies.
Dr. David has been involved for nearly 25 years, mostly with Gernot Vogel, in systematic research on several
groups of Asian reptiles, especially the Trimeresurus-complex and the genera Oligodon, Amphiesma, Xenochrophis, and Cyrtodactylus. His geographic areas of interest include India, Thailand, China, Vietnam, and
especially Laos and Sumatra (Indonesia). Dr. David has also addressed problems of nomenclature affecting
various taxa of snakes and lizards. As of July 2014, Dr. David is the author or co-author of 121 publications,
including five monographs or books. He has co-authored the description of a total of 31 new species of amphibians and reptiles. Along with Gernot Vogel, Dr. David is preparing a monograph on Asian pitvipers and
the snake fauna of Sumatra.
Irvan Sidik was born in Bandung, West Java Province, Indonesia. Irvan obtained a Masters of Science degree
in the field of phylogenetics at the Institute Technology of Bandung. Since 1992 Irvan has worked as a staff
researcher in the laboratory of herpetology at the Museum Zoologicum Bogoriense, Indonesian Institute of
Sciences (LIPI) in the Cibinong Science Center. Beginning as an auxiliary field survey researcher, and then
as a local CITES officer, Irvan became interested and developed a great interest in the snakes of the region
of Sundaland. Irvan has continued with more scholarly work on the mountainous areas of the western part
of Indonesia. Irvans research is based on museum collections of specimens and field research in Indonesias
regions mentioned above. Irvan has been involved in several international research collaborations, and is currently working with the University of Texas at Arlington, USA on research of amphibians and reptiles in the
mountains of Java and Sumatra. Irvan has published on the herpetofauna of Kalimantan and his first book was
about snakes that are traded in Indonesia (CITES appendices I, II, and III) written in Indonesian. Currently,
Irvan is studying the phylogeography of the reed snake genera Calamaria for his Ph.D. at the University of
Brawijaya, Malang.
.
In accordance with the International Code of Zoological Nomenclature new rules and regulations (ICZN 2012), we have deposited this paper in publicly accessible institutional libraries.
The new species described herein has been registered in ZooBank (Polaszek 2005a, b), the official online registration system for the ICZN. The ZooBank publication LSID (Life Science
Identifier) for the new species described here can be viewed through any standard web browser by appending the LSID to the prefix http://zoobank.org/. The LSID for this publication
is: urn:lsid:zoobank.org:pub:27336534-BAFC-40BE-84F7-43E0334596CD.
Separate print-only edition of paper(s) (reprint) are available upon request as a print-on-demand service. Please inquire by sending a request to: Amphibian & Reptile Conservation
(amphibian-reptile-conservation.org; arc.publisher@gmail.com).
Amphibian & Reptile Conservation is a Content Partner with the Encyclopedia of Life (EOL); http:///www.eol.org/ and submits information about new species to the EOL freely.
Digital archiving of this paper are found at the following institutions: ZenScientist (http://www.zenscientist.com/index.php/filedrawer); Ernst Mayr Library, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts (USA); Florida Museum of Natural History, Gainesville, Florida (USA).
Complete journal archiving is found at: ZenScientist (http://www.zenscientist.com/index.php/filedrawer); Florida Museum of Natural History, Gainesville, Florida (USA).
Citations
ICZN. 2012. Amendment of Articles 8,9,10,21 and 78 of the International Code of Zoological Nomenclature to expand and refine methods of publication. Zootaxa 3450: 17.
Polaszek A et al. 2005a. Commentary: A universal register for animal names. Nature 437: 477.
Polaszek A et al. 2005b. ZooBank: The open-access register for zoological taxonomy: Technical Discussion Paper. Bulletin of Zoological Nomenclature 62(4): 210220.